Ulvophyceae
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The Ulvophyceae or ulvophytes are a class of
green algae The green algae (: green alga) are a group of chlorophyll-containing autotrophic eukaryotes consisting of the phylum Prasinodermophyta and its unnamed sister group that contains the Chlorophyta and Charophyta/ Streptophyta. The land plants ...
, distinguished mainly on the basis of ultrastructural morphology, life cycle and
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
data. The sea lettuce, ''Ulva'', belongs here. Other well-known members include '' Caulerpa'', '' Codium'', ''
Acetabularia ''Acetabularia'' is a genus of green algae in the family Polyphysaceae. Typically found in subtropical waters, ''Acetabularia'' is a single-celled organism, but gigantic in size and complex in form, making it an excellent model organism for stud ...
'', '' Cladophora'', '' Trentepohlia'' and '' Monostroma''. The Ulvophytes are diverse in their
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
and their
habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...
. Most are
seaweed Seaweed, or macroalgae, refers to thousands of species of macroscopic, multicellular, marine algae. The term includes some types of ''Rhodophyta'' (red), '' Phaeophyta'' (brown) and ''Chlorophyta'' (green) macroalgae. Seaweed species such as ...
s such as those listed above. Others, such as '' Rhizoclonium'', '' Pithophora'' and some species of '' Cladophora'' live in fresh water and in some areas are considered
weed A weed is a plant considered undesirable in a particular situation, growing where it conflicts with human preferences, needs, or goals.Harlan, J. R., & deWet, J. M. (1965). Some thoughts about weeds. ''Economic botany'', ''19''(1), 16-24. Pla ...
s.


Morphology

Ulvophycean algae are diverse in morphology. The thalli are typically colonial (some are unicellular). A few taxa have
flagella A flagellum (; : flagella) (Latin for 'whip' or 'scourge') is a hair-like appendage that protrudes from certain plant and animal sperm cells, from fungal spores ( zoospores), and from a wide range of microorganisms to provide motility. Many pr ...
in their vegetative stage, such as '' Oltmannsiellopsis''. Common forms include filaments (both unbranched and branched), tubular and blade-like thalli (such as in ''
Ulva Ulva (; ) is a small island in the Inner Hebrides of Scotland, off the west coast of Mull. It is separated from Mull by a narrow strait, and connected to the neighbouring island of Gometra by a bridge. Much of the island is formed from Cen ...
''), and siphonous thalli. Siphonous thalli are composed of a single giant cell (siphon) with thousands of nuclei and chloroplasts. The life cycle of members of the Ulvophyceae are similarly diverse. Species often reproduce asexually by forming aplanospores,
akinete An akinete is an enveloped, thick-walled, non-motile, dormant cell (biology), cell formed by both cyanobacteria and algae. Cyanobacterial akinetes are mainly formed by filamentous, heterocyst-forming members under the order Nostocales and Stigone ...
s or
zoospore A zoospore is a motile asexual spore that uses a flagellum for locomotion in aqueous or moist environments. Also called a swarm spore, these spores are created by some protists, bacteria, and fungi to propagate themselves. Certain zoospores are ...
s; zoospores have two or four flagella. Asexual reproduction may also occur through fragmentation of the thallus, where the pieces of the thallus grow into a new organism.
Sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...
is isogamous or anisogamous, and may be haplontic or diplohaplontic (i.e., involving an alternation of generations). In several taxa, the zygote exists as a small, microscopic dormant stage (also interpreted as a unicellular
sporophyte A sporophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the biological life cycle, life cycles of plants and algae. It is a diploid multicellular organism which produces asexual Spo ...
) and is called the '' Codiolum'' phase.


Ultrastructure

The class Ulvophyceae was originally defined by K. R. Mattox and K. D. Stewart based on ultrastructural characters. These were: a counter-clockwise arrangement of the
flagella A flagellum (; : flagella) (Latin for 'whip' or 'scourge') is a hair-like appendage that protrudes from certain plant and animal sperm cells, from fungal spores ( zoospores), and from a wide range of microorganisms to provide motility. Many pr ...
r
basal bodies A basal body (synonymous with basal granule, kinetosome, and in older cytological literature with blepharoplast) is a protein structure found at the base of a eukaryotic undulipodium (cilium or flagellum). The basal body was named by Theodor ...
,
cytokinesis Cytokinesis () is the part of the cell division process and part of mitosis during which the cytoplasm of a single eukaryotic cell divides into two daughter cells. Cytoplasmic division begins during or after the late stages of nuclear division ...
by furrowing, a closed persistent
mitotic spindle In cell biology, the spindle apparatus is the cytoskeletal structure of eukaryotic cells that forms during cell division to separate sister chromatids between daughter cells. It is referred to as the mitotic spindle during mitosis, a process ...
, and the lack of a phycoplast. The
cell wall A cell wall is a structural layer that surrounds some Cell type, cell types, found immediately outside the cell membrane. It can be tough, flexible, and sometimes rigid. Primarily, it provides the cell with structural support, shape, protection, ...
composition and structure are highly variable, and in the flagellate life stages of some organisms, there is no cell wall altogether. The order
Trentepohliales Trentepohliaceae are a family (biology), family of green algae, and the only family in the order Trentepohliales. It is a member of the Ulvophyceaen green algae. The family is characterized by traits like net-like chloroplasts without pyrenoids, ...
has a unique set of ultrastructural features (such as the presence of a phragmoplast), and was once considered to be its own separate class, the Trentepohliophyceae. Most of these ultrastructural characters are found in other lineages of algae, which has made defining Ulvophyceae difficult.


Ecology

The Ulvophyceae are most diverse in marine habitats. A substantial number of species also live in
brackish Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estuari ...
or
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. The term excludes seawater and brackish water, but it does include non-salty mi ...
habitats; in temperate regions they are typically most abundant during the spring and summer months. They may be found in still (e.g. lakes and ponds) or flowing waters (e.g. rivers and streams); they are typically attached to a substrate such as rocks, sediments, other aquatic algae, or occasionally animals like
turtle Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
s or
mollusc Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
shells. Similarly, some species live in terrestrial habitats growing on soil, rocks, trees, plants, and other artificial substrates. Ulvophycean algae are often beneficial to their habitats; for example, '' Cladophora'' has been described as an
ecosystem engineer An ecosystem engineer is any species that creates, significantly modifies, maintains or destroys a habitat. These organisms can have a large impact on species richness and landscape-level heterogeneity of an area. As a result, ecosystem engine ...
and provides a range of niches for other aquatic organisms. However, ''Cladophora'' and a few other taxa, such as '' Pithophora'' and ''
Ulva Ulva (; ) is a small island in the Inner Hebrides of Scotland, off the west coast of Mull. It is separated from Mull by a narrow strait, and connected to the neighbouring island of Gometra by a bridge. Much of the island is formed from Cen ...
'' often form nuisance blooms due to
eutrophication Eutrophication is a general term describing a process in which nutrients accumulate in a body of water, resulting in an increased growth of organisms that may deplete the oxygen in the water; ie. the process of too many plants growing on the s ...
. Some species of ulvophytes are
endosymbiotic An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualistic relationship. Examples are nitrogen-fixing bacteria (called rhizobia), which live in the root ...
within other algae, such as '' Ulvella'' which grows within other filamentous algae. Within the terrestrial order Trentepohliales, some taxa form symbioses with
lichen A lichen ( , ) is a hybrid colony (biology), colony of algae or cyanobacteria living symbiotically among hypha, filaments of multiple fungus species, along with yeasts and bacteria embedded in the cortex or "skin", in a mutualism (biology), m ...
s as their phycobionts. Marine lichens also host some Ulvophyceae as symbionts; the diversity of these symbiotic algae is currently under-studied. The family Trentepohliaceae is unusual in that it contains a few
parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The ent ...
genera. For example, '' Cephaleuros'' is an endophyte within the leaves of
vascular plant Vascular plants (), also called tracheophytes (, ) or collectively tracheophyta (; ), are plants that have lignin, lignified tissues (the xylem) for conducting water and minerals throughout the plant. They also have a specialized non-lignified Ti ...
s. ''Cephaleuros'' can cause significant economic damage to crop plants.


Evolution

The origin and early diversification of the Ulvophyceae likely took place in the late
Neoproterozoic The Neoproterozoic Era is the last of the three geologic eras of the Proterozoic geologic eon, eon, spanning from 1 billion to 538.8 million years ago, and is the last era of the Precambrian "supereon". It is preceded by the Mesoproterozoic era an ...
, though may have taken place earlier, in the Mesoproterozoic. Although most contemporary ulvophytes are marine macroalgae (
seaweed Seaweed, or macroalgae, refers to thousands of species of macroscopic, multicellular, marine algae. The term includes some types of ''Rhodophyta'' (red), '' Phaeophyta'' (brown) and ''Chlorophyta'' (green) macroalgae. Seaweed species such as ...
s), ancestral ulvophytes may have been freshwater, unicellular green algae. Molecular phylogenetic evidence suggests that macroscopic growth was achieved independently in the various major lineages of Ulvophyceae ( Ulvales-
Ulotrichales Ulotrichales is an order of green algae in the class Ulvophyceae.See the NCBIbr>webpage on Ulotrichales Data extracted from the Genera unplaced to family (''incertae sedis''): *'' Trichosarcina'' Families * '' Binucleariaceae'' * '' Collin ...
,
Trentepohliales Trentepohliaceae are a family (biology), family of green algae, and the only family in the order Trentepohliales. It is a member of the Ulvophyceaen green algae. The family is characterized by traits like net-like chloroplasts without pyrenoids, ...
, Cladophorales, Bryopsidales and Dasycladales). The
fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
of Ulvophyceae is generally sparse, as most algae do not
biomineralize Biomineralization, also written biomineralisation, is the process by which living organisms produce minerals, often resulting in hardened or stiffened '' mineralized tissues''. It is an extremely widespread phenomenon: all six taxonomic kingd ...
well. Nevertheless, some orders such as the Dasycladales and Bryopsidales have calcified thalli and a more extensive fossil record. The earliest known representative is the Cladophorales ''Proterocladus antiquus'' from the
Mesoproterozoic The Mesoproterozoic Era is a geologic era that occurred from . The Mesoproterozoic was the first era of Earth's history for which a fairly definitive geological record survives. Continents existed during the preceding era (the Paleoproterozoic ...
-
Neoproterozoic The Neoproterozoic Era is the last of the three geologic eras of the Proterozoic geologic eon, eon, spanning from 1 billion to 538.8 million years ago, and is the last era of the Precambrian "supereon". It is preceded by the Mesoproterozoic era an ...
boundary in North China. However, its taxonomic placement is not certain. Because Ulvophyceae lacks a consistent set of ultrastructural characteristics, its
monophyly In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent comm ...
has been questioned.
Molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
studies have found Ulvophyceae to be monophyletic, albeit without strong support. The internal relationships between the different clades is also unclear. Current hypothesis on relationships among the main clades of Ulvophyceae are shown below:


See also

* List of Ulvophyceae genera


References

{{Taxonbar, from=Q132016 Green algae classes